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Plumage () is a layer of
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and an exa ...
s that covers a
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage differ between
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
and subspecies and may vary with age classes. Within species, there can be different colour morphs. The placement of feathers on a bird is not haphazard but rather emerges in organized, overlapping rows and groups, and these are known by standardized names. Most birds
moult In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is a process by which an animal casts off parts of its body to serve some beneficial purpose, either at ...
twice a year, resulting in a breeding or ''nuptial plumage'' and a ''basic plumage''. Many
duck Duck is the common name for numerous species of waterfowl in the family (biology), family Anatidae. Ducks are generally smaller and shorter-necked than swans and goose, geese, which are members of the same family. Divided among several subfam ...
s and some other species such as the
red junglefowl The red junglefowl (''Gallus gallus''), also known as the Indian red junglefowl (and formerly the bankiva or bankiva-fowl), is a species A species () is often defined as the largest group of organisms in which any two individuals of the a ...
have males wearing a bright nuptial plumage while breeding and a drab ''eclipse plumage'' for some months afterward. The painted bunting's juveniles have two inserted moults in their first autumn, each yielding plumage like an adult female. The first starts a few days after fledging replacing the ''juvenile plumage'' with an ''auxiliary formative plumage''; the second a month or so later giving the ''formative plumage''. Abnormal plumages include a variety of conditions.
Albinism Albinism is the congenital absence of melanin in an animal or plant resulting in white hair, feathers, scales and skin and reddish pink or blue eyes. Individuals with the condition are referred to as albinos. Varied use and interpretation of ...
, total loss of colour, is rare, but partial loss of colours is more common. Some species are colour polymorphic, having two or more colour variants. A few species have special types of polymorphism, as in the male
ruff Ruff may refer to: Places *Ruff, Virginia, United States, an unincorporated community *Ruff, Washington, United States, an unincorporated community Other uses *Ruff (bird) (''Calidris pugnax'' or ''Philomachus pugnax''), a bird in the wader famil ...
which has an assortment of different colours around the head and neck in the breeding season only. Hen feathering is an inherited plumage character in domestic fowl controlled by a single gene. '' Plumology'' (or ''plumage science'') is the name for the science that is associated with the study of feathers.


Humphrey–Parkes (H–P) moult and plumage terminology

Almost all species of birds
moult In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is a process by which an animal casts off parts of its body to serve some beneficial purpose, either at ...
at least annually, usually after the breeding season, known as the ''pre-basic moult''. This resulting covering of feathers, which will last either until the next breeding season or until the next annual moult, is known as the basic plumage. Many species undertake another moult before the breeding season known as the ''pre-alternate moult'', the resulting breeding plumage being known as the ''alternate plumage'' or nuptial plumage. The alternate plumage is often brighter than the basic plumage, for sexual display, but may also be cryptic to hide incubating birds that might be vulnerable on the nest. The
Humphrey–Parkes terminology Humphrey–Parkes terminology is a system of nomenclature for the plumage of birds. Before the Humphrey–Parkes system, plumage was named after the belief that a certain plumage was breeding plumage and others were not. However, as this system did ...
requires some attention to detail to name moults and plumages correctly.


Eclipse plumage

Many male ducks have bright, colourful plumage, exhibiting strong
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
. However, they moult into a dull plumage after breeding in mid-summer. This drab, female-like appearance is called ''eclipse plumage''. When they shed feathers to go into an eclipse, the ducks become flightless for a short period. Some duck species remain in eclipse for one to three months in the late summer and early fall, while others retain the cryptic plumage until the next spring when they undergo another moult to return to their breeding plumage. Although mainly found in the
Anatidae The Anatidae are the biological family (biology), family of water birds that includes ducks, goose, geese, and swans. The family has a cosmopolitan distribution, occurring on all the world's continents except Antarctica. These birds are adapted f ...
, a few other species, including related
red junglefowl The red junglefowl (''Gallus gallus''), also known as the Indian red junglefowl (and formerly the bankiva or bankiva-fowl), is a species A species () is often defined as the largest group of organisms in which any two individuals of the a ...
, most fairywrens and some
sunbird Sunbirds and spiderhunters make up the family Nectariniidae of passerine birds. They are small, slender passerines from the Old World, usually with downward-curved bills. Many are brightly coloured, often with iridescent feathers, particularly i ...
s also have an eclipse plumage. In the superb and splendid fairywrens, very old males (over about four years) may moult from one nuptial plumage to another whereas in the red-backed and white-winged fairywrens, males do not acquire nuptial plumage until four years of age – well after they become sexually mature and indeed longer than the vast majority of individuals live. In contrast to the ducks, males of hummingbirds and most lek-mating passerines – like the Guianan cock-of-the-rock or birds of paradise – retain their exuberant plumage and sexual dimorphism at all times, moulting as ordinary birds do once annually.


Abnormal plumages

There are hereditary as well as non-hereditary variations in plumage that are rare and termed abnormal or aberrant plumages. Melanism refers to an excess of black or dark colours. Erythromelanism or
erythrism Erythrism or erythrochroism refers to an unusual reddish pigmentation of an animal's hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, ap ...
is the result of excessive reddish-brown erythromelanin deposition in feathers that normally lack melanin. Melanin of different forms combine with xanthophylls to produce colour mixtures and when this combination is imbalanced it produces colour shifts that are termed schizochroisms (including xanthochromism – an overabundance of yellow – and axanthism – lack of yellow – which are commonly bred in
cagebird Aviculture is the practice of keeping and breeding birds, especially of wild birds in captivity. Aviculture Aviculture is the practice of keeping birds (class ''Aves'') in captivity in controlled conditions, normally within the confines of a c ...
s such as budgerigars). A reduction in eumelanin leads to non-eumelanin schizochroism with an overall fawn plumage while a lack of phaeomelanin results in grey-coloured non-phaeomelanin schizochroism. Carotenism refers to the abnormal distribution of
carotenoid Carotenoids () are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, archaea, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, corn, tomatoes, cana ...
pigments. The term "dilution" is used for situations where the colour is of a lower intensity overall; it is caused by decreased deposition of pigment in the developing feather, and can thus not occur in
structural coloration Structural coloration in animals, and a few plants, is the production of colour by microscopically structured surfaces fine enough to interfere with visible light instead of Biological pigment, pigments, although some structural coloration occu ...
(i.e., "dilute blue" does not exist); pale structural colors are instead achieved by shifting the peak wavelength at which light is refracted. Dilution regularly occurs in normal plumage (grey, buff, pink and cream colours are usually produced by this process), but may in addition occur as an aberration (e.g., all normally black plumage becoming grey). In some birds – many
true owl The true owls or typical owls (family (biology), family Strigidae) are one of the two generally accepted families of owls, the other being the barn owls and bay owls (Tytonidae). This large family comprises 230 living or recently extinct species ...
s (Strigidae), some
nightjar Nightjars are medium-sized nocturnal or crepuscular birds in the family Caprimulgidae and order Caprimulgiformes, characterised by long wings, short legs, and very short bills. They are sometimes called bugeaters, their primary source of food ...
s (Caprimulgidae) and a few
cuckoo Cuckoos are birds in the Cuculidae ( ) family, the sole taxon in the order Cuculiformes ( ). The cuckoo family includes the common or European cuckoo, roadrunners, koels, malkohas, couas, coucals, and anis. The coucals and anis are somet ...
s (''
Cuculus ''Cuculus'' is a genus of cuckoos which has representatives in most of the Old World, although the greatest diversity is in tropical southern and southeastern Asia. Taxonomy The genus ''Cuculus'' was introduced in 1758 by the Swedish naturalist ...
'' and relatives) being widely known examples – there is colour polymorphism. This means that two or more colour variants are numerous within their
population Population is a set of humans or other organisms in a given region or area. Governments conduct a census to quantify the resident population size within a given jurisdiction. The term is also applied to non-human animals, microorganisms, and pl ...
s during all or at least most seasons and plumages; in the above-mentioned examples a brown (phaeomelanin) and grey (eumelanin) morph exist, termed "hepatic form" particularly in the cuckoos. Other cases of natural polymorphism are of various kinds; many are melanic/nonmelanic (some paradise-flycatchers, ''Terpsiphone'', for example), but more unusual types of polymorphism exist – the face colour of the Gouldian finch (''Erythrura' gouldiae'') or the courtship types of male
ruff Ruff may refer to: Places *Ruff, Virginia, United States, an unincorporated community *Ruff, Washington, United States, an unincorporated community Other uses *Ruff (bird) (''Calidris pugnax'' or ''Philomachus pugnax''), a bird in the wader famil ...
s (''Philomachus pugnax'').


Albinism

Albinism Albinism is the congenital absence of melanin in an animal or plant resulting in white hair, feathers, scales and skin and reddish pink or blue eyes. Individuals with the condition are referred to as albinos. Varied use and interpretation of ...
in birds is rare, occurring to any extent in perhaps one in 1800 individuals. It involves loss of colour in all parts including the iris of the eyes, bills, skin, legs, and feet. It is usually the result of a genetic mutation causing the absence of
tyrosinase Tyrosinase is an oxidase that is the rate-limiting enzyme for controlling the production of melanin. The enzyme is mainly involved in two distinct reactions of melanin synthesis otherwise known as the Raper–Mason pathway. Firstly, the hydroxy ...
, an enzyme essential for melanin synthesis.
Leucism Leucism () is a wide variety of conditions that result in partial loss of pigmentation in an animal—causing white, pale, or patchy coloration of the skin, hair, feathers, scales, or cuticles, but not the eyes. It is occasionally spelled ''le ...
(which includes what used to be termed as "partial albinism") refers to loss of pigments in some or all parts of feathers. A bird that is
albino Albinism is the congenital absence of melanin in an animal or plant resulting in white hair, feathers, scales and skin and reddish pink or blue eyes. Individuals with the condition are referred to as albinos. Varied use and interpretation of ...
(from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
''albus'', "white") has white
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and an exa ...
s in place of coloured ones on some portion of its body. A bird that is naturally white, such as a
swan Swans are birds of the genus ''Cygnus'' within the family Anatidae. The swans' closest relatives include the goose, geese and ducks. Swans are grouped with the closely related geese in the subfamily Anserinae where they form the tribe (biology) ...
,
goose A goose (: geese) is a bird of any of several waterfowl species in the family Anatidae. This group comprises the genera '' Anser'' (grey geese and white geese) and '' Branta'' (black geese). Some members of the Tadorninae subfamily (e.g., Egy ...
, or
egret Egrets ( ) are herons, generally long-legged wading birds, that have white or buff plumage, developing fine plumes (usually milky white) during the breeding season. Egrets are not a biologically distinct group from herons and have the same build ...
, is not an albino, nor is a bird that has seasonally alternating white plumage."Albinism". Terres, John. K. ''The Audubon Society Encyclopedia of North American Birds'', New York: Alfred A. Knopf, 1980. . Four degrees of albinism have been described. The most common form is termed ''partial albinism'', in which local areas of the bird's body, such as certain feathers, are lacking the pigment
melanin Melanin (; ) is a family of biomolecules organized as oligomers or polymers, which among other functions provide the pigments of many organisms. Melanin pigments are produced in a specialized group of cells known as melanocytes. There are ...
. The white areas may be symmetrical, with both sides of the bird showing a similar pattern. In ''imperfect albinism'', the pigment is partially inhibited in the
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other animal coverings, such as the arthropod exoskeleton, have different ...
,
eye An eye is a sensory organ that allows an organism to perceive visual information. It detects light and converts it into electro-chemical impulses in neurons (neurones). It is part of an organism's visual system. In higher organisms, the ey ...
s, or feathers, but is not absent from any of them. ''Incomplete albinism'' is the complete absence of pigment from the skin, eyes, or feathers, but not all three. A completely albino bird is the most rare. The eyes in this case are pink or red, because
blood Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood is com ...
shows through in the absence of pigment in the irises. The
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for pecking, grasping, and holding (in probing for food, eating, manipulating and ...
, legs, and feet are very pale or white. Albino adults are rare in the wild because their eyesight is poor resulting in greater risk of predation. They are likely easier targets for
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
s because their colour distinguishes them from their environment. Falconers have observed that their trained birds are likely to attack a white
pigeon Columbidae is a bird family consisting of doves and pigeons. It is the only family in the order Columbiformes. These are stout-bodied birds with small heads, relatively short necks and slender bills that in some species feature fleshy ceres. ...
in a flock because it is conspicuous. A complete albino often has weak eyesight and brittle wing and tail feathers, which may reduce its ability to fly. In flocks, albinos are often harassed by their own species. Such observations have been made among red-winged blackbirds,
barn swallow The barn swallow (''Hirundo rustica'') is the most widespread species of swallow in the world, occurring on all continents, with vagrants reported even in Antarctica. It is a distinctive passerine bird with blue upperparts and a long, deeply f ...
s, and
African penguin The African penguin (''Spheniscus demersus''), also known as Cape penguin or South African penguin, is a species of penguin confined to southern African waters. It is the only penguin found in the Old World. Like all penguins, it is flightless, ...
s. In a nesting colony of the latter, three unusual juveniles—one black-headed, one white-headed, and one full albino—were shunned and abused by companions. Albinism has been reported in all
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
s and in 54
families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...
of North American birds. The
American robin The American robin (''Turdus migratorius'') is a migratory bird of the true thrush genus and Turdidae, the wider thrush family. It is named after the European robin because of its reddish-orange breast, though the two species are not clos ...
and
house sparrow The house sparrow (''Passer domesticus'') is a bird of the Old World sparrow, sparrow family Passeridae, found in most parts of the world. It is a small bird that has a typical length of and a mass of . Females and young birds are coloured pa ...
led bird species in the incidence of albinism. Albinistic white appears to replace brown pigments more often than red or yellow ones; records suggest a greater incidence in
crow A crow is a bird of the genus ''Corvus'', or more broadly, a synonym for all of ''Corvus''. The word "crow" is used as part of the common name of many species. The related term "raven" is not linked scientifically to any certain trait but is rathe ...
s,
raven A raven is any of several large-bodied passerine bird species in the genus '' Corvus''. These species do not form a single taxonomic group within the genus. There is no consistent distinction between crows and ravens; the two names are assigne ...
s, and
hawk Hawks are birds of prey of the family Accipitridae. They are very widely distributed and are found on all continents, except Antarctica. The subfamily Accipitrinae includes goshawks, sparrowhawks, sharp-shinned hawks, and others. This ...
s than in goldfinches or orioles. Several kinds of albinism in chickens has been described: A complete albinism controlled by an autosomal recessive gene and two different kinds of partial albinism. One of the partial albinisms is sex-linked and the other is autosomal recessive. A fourth kind of albinism severely reduce pigmentation in the eyes, but only dilutes the pigment in the plumage. Abnormally white feathers are not always due to albinism. Injury or disease may change their color, including dietary deficiencies or circulatory problems during feather development. Aging may also turn a bird's feathers white.


Hen feathering in cocks

Hen feathering in cocks is a genetically conditioned character in domestic fowl ( Gallus gallus domesticus). Males with this condition develop a female-type plumage, although otherwise look and respond like virile males. In some breeds, one can see males that have a plumage completely similar in all aspects to that of females. The trait is controlled by a simple autosomic
dominant gene In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and ...
, whose expression is limited to the male sex. The condition is due to an enhanced activity of the aromatase complex of
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
s responsible for
estrogen Estrogen (also spelled oestrogen in British English; see spelling differences) is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three ...
synthesis. So estrogen formation in the skin is as much as several hundred-fold higher than that of normal chickens.


Pigmentation conditions

*
Albinism Albinism is the congenital absence of melanin in an animal or plant resulting in white hair, feathers, scales and skin and reddish pink or blue eyes. Individuals with the condition are referred to as albinos. Varied use and interpretation of ...
, the lack of melanin pigmentation *
Leucism Leucism () is a wide variety of conditions that result in partial loss of pigmentation in an animal—causing white, pale, or patchy coloration of the skin, hair, feathers, scales, or cuticles, but not the eyes. It is occasionally spelled ''le ...
, a condition similar to albinism in animals, characterized by reduced pigmentation in general *
Melanism Melanism is the congenital excess of melanin in an organism resulting in dark pigment. Pseudomelanism, also called abundism, is another variant of pigmentation, identifiable by dark spots or enlarged stripes, which cover a large part of the bod ...
(or melanosis), unusually dark melanin pigmentation * Xanthochromism, unusually yellow pigmentation * Ino budgerigar mutation, the occurrence of this mutation in captive-bred
budgerigar The budgerigar ( ; ''Melopsittacus undulatus''), also known as the common parakeet, shell parakeet or budgie ( ), is a small, long-tailed, seed-eating parrot native to Australia. Naturally the species is green and yellow with black, scallop ...
s * Axanthism, lack of yellow pigmentation


See also

*
Animal coloration Animal coloration is the general appearance of an animal resulting from the reflection or emission of light from its surfaces. Some animals are brightly coloured, while others are hard to see. In some species, such as the peafowl, the male h ...
* Hen feathering * Imping * Pluma porgy *
Plume hunting Plume hunting is the hunting of wild birds to harvest their feathers, especially the more decorative plumes which were sold for use as ornamentation, particularly in hat-making (millinery). The movement against the plume trade in the United K ...
*
Sea pen Sea pens are marine cnidarians belonging to the superfamily Pennatuloidea, which are colony-forming benthic filter feeders within the order Scleralcyonacea. There are 14 families within the order and 35 extant genera, and it is estimated a ...
*
Vermiculation Vermiculation is a surface pattern of dense but irregular lines, so called from the Latin meaning "little worm" because the shapes resemble worms, worm casts, or worm tracks in mud or wet sand. The word may be used in a number of contexts for ...


Notes

Males of the white-shouldered and emperor fairywrens of
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...
do not enter an eclipse plumage.


References

{{Birds Bird anatomy Feathers